Argulus ambystoma, a New Parasitic on the Salamander Ambystoma dumerilii from Mexico (Crustacea: : )1

WILLIAM J. POLY2, Department of Zoology, Southern Illinois University, Carbondale, IL 62901-6501

ABSTRACT. A new species of is described based on 18 specimens taken from the salamander ("achoque" or "ajolote") Ambystoma dumerilii Duges, collected in Lake Patzcuaro, Michoacan, Mexico. Diagnostic characters include the shape of the respiratory areas, number of sclerites in suction cup rods, and structures on the legs of males. Females are heavily stippled, whereas males have a very distinctive pigment pattern consisting of abundant melanophores covering the testes dors ally and two dark, inverted triangular patches on the carapace dorsally. The new species is similar to the North American species, A versicolor,A. americanus, A. maculosus, and A diversus. A single, dorsal pore was observed on each caudal ramus using scanning electron microscopy; these pores have not been reported previously in the Branchiura.

OHIO J SCI 103 (3>52-6l, 2003

INTRODUCTION 1991; Osorio-Sarabia and others 1986; Perez-Ponce de Ten species of Argulus Miiller have been collected in Leon and others 1994; Peresbarbosa-Rojas and others Mexico, Central America, and the West Indies. The wide- 1994; Espinosa-Huerta and others 1996; Peresbarbosa- spread exotic, Argulus japonicus Thiele, was found on Rojas and others 1997). A number of endemic taxa exist aquarium fishes, Carassius auratus (Linne) and Astro- in Lake Patzcuaro including several fishes (Cbirostoma notus ocellatus (Agassiz), in Puerto Rico (Bunkley-Williams patzcuaro Meek, C. e. estor Jordan, C. a. attenuatum and Williams 1994), and Vargas and Fallas (1976) found Meek), a crayfish (Cambarellus patzcuarensis Villa- A. japonicus on Carassius sp. in Costa Rica in 1972 and lobos), and the salamander host of the new argulid (Am- 1973- Argulus dactylopteri Thorell was described from bystoma dumerilii (Duges)) (Brandon 1970; Barbour specimens collected from the gills of Dactylopterus 1973; Hobbs 1989). volitans (Linne) in the West Indies (Thorell 1865, 1867); neither Cressey (1982) nor Pineda and others (1995) MATERIALS AND METHODS mentioned A. dactylopteri. Wilson (1902) gave the locality Three males and two females were examined using for A. dactylopteri as the East Indian Ocean from the scanning electron microscopy (SEM). Preparation branchial cavity of D. volitans, whereas Yamaguti (1963) procedures were modified slightly from those of Rupp cited Europe and Dactylopterus sp. If the type locality (1990). All specimens were initially preserved and stored of A. dactylopteri is somewhere other than the Atlantic in 70% ethanol (EtOH, J. A. Beatty pers. comm.). Speci- Ocean or Mediterranean Sea, the host would not be D. mens were dehydrated in an EtOH series consisting of volitans but a species of Dactyloptena (see Eschmeyer 80% (5 min), 90% (5 min), 100% (1st, 5 min; 2nd, 10 min), 1997). Only six species of Argulus, A. chromidis Kr0yer; then immediately critical point dried (CO2), mounted A. flavescens Wilson; A. megalops Smith; A. mexicanus on metal stubs with carbon paint, allowed to dry over- Pineda, Paramo, and Del Rio; A. melanostictus Wilson; night in an oven (60° C), and sputter coated with gold/ and A. rhamdiae Wilson have been found in Mexico palladium. Specimens were stored in an oven at 60° C (Wilson 1936; Causey I960; Olson 1972; Cressey 1982; between uses. Some specimens were coated up to three Fucugauchi and others 1988; Pineda and others 1995; times and were examined in a Hitachi S570 Scanning Suarez-Morales and Gasca 1997; Suarez-Morales and Electron Microscope at 15kV. Six females and seven others 1998). In addition, an unidentified species of Argu- males (including the holotype, all adult) were examined lus was reported from Potamarius nelsoni (Evermann under dissecting and compound microscopes in a and Goldsborough) by Pineda-Lopez and others (1985). watchglass or temporary slide mount (with 70% EtOH/ The species described herein was collected from glycerin). The holotype and allotype were photographed Lake Patzcuaro (Lago de Patzcuaro) in Central Mexico, using a camera tube on a light microscope. In addition, State of Michoacan. Both the invertebrate and vertebrate some counts and measurements were taken from 3 male faunas of the lake have been investigated; however, the and 2 female specimens later used for SEM. All Argulus has not been recorded as a component measurements were made using an ocular micrometer, of the fauna (Ueno 1939; Ancona and others 1940; and measurements reported in the description are Brehm 1942; Tressler 1954; Brandon 1970; Barbour arranged as follows: range (mean, holotype) with allo- 1973; Rosas and others 1985; Chacon-Torres and others type values substituted in the case of females. Values for the right and left sides refer to the right and left of the specimen in dorsal view. Type specimens were deposited 'Manuscript received 20 September 2001 and in revised form 5 in the National Museum of Natural History, Washington, May 2002 (#01-24). DC (USNM), and specimens of Argulus americanus 2Present Address: Department of Ichthyology, California Academy of Sciences, Golden Gate Park, San Francisco, CA 94118 Wilson (USNM 274331, 274332), A. diversusWihon (USNM OHIO JOURNAL OF SCIENCE W. J. POLY 53

32823,32824), A. maculosusWilsonOJSNM 12226,74329), and A. versicolor Wilson (USNM 74321; MCZ 19604, 19607) were examined by light microscopy for comparative purposes (MCZ, Museum of Comparative Zoology, Harvard University, Cambridge, MA).

RESULTS Family Argulidae Rafinesque, 1815 Argulus Miiller, 1785 Argulus ambystoma, new species

Material Examined Holotype: adult male, 3-48 mm total length, USNM 282782, Lake Patzcuaro, Michoacan, Mexico. Allotype: adult female, 4.18 mm total length, USNM 282783, col- lected with holotype. Paratypes: 5 adult males, 4 adult females, USNM 282784, collected with holotype; 4 adult males, 3 adult females (used for SEM or cleared, not FIGURE 1. Argulus ambystoma, new species, dorsal view, male (holo- retained), collected with holotype (host collected by type, left) and female (allotype, right) (scale bar = 1.0 mm). J. Arnett; Argulus collected from hosts, Ambystoma dumerilii (Duges), byj. A. Beatty and R. A. Brandon on 27 May 1981). scattered on dorsal surface and margins of carapace. Smaller type of sensillum present on anterior margin Diagnosis (between sinuses) of cephalic region, abundant in inter- Respiratory areas with a smaller, oblong "area" sit- spaces between larger sensilla. Pair of compound eyes uated anterior to and abutting or nearly contacting a anteriorly with diameters (left and right eyes, Jim) 170- larger posterior "area"; respiratory areas outlined with 210 (187, left: 170 and right: 180) (n = 18 eyes) in males, melanophores and very distinct; caudal rami near pos- 200-240 (224, left: 230 and right: 210) (n = 14 eyes) in terior end of abdomen in anal sinus; paired post- females. Transverse distance between eyes (|Xm) 390- antennal spines with reduced anterior spine and stout 440 (419, 410) in males, 510-650 (604, 570) in females. posterior spine; males with 43-56 support rods per suc- Nauplius eye with one anterior and two posterior ocelli, tion cup; females with 45-54 support rods per suction pigment between ocelli forming a dark, prominent "Y" cup; usually 3 to 4 sclerites per rod in males and 4 to 5 or "V." Sclerotized dorsal ridges forked anterior of eyes, sclerites per rod in females; basal plate of second maxilla inner branches longer and less developed than outer with three stout, digitate spines; first two pairs of legs branches. Bifurcation of dorsal ridges usually more ob- each with a recurved flagellum on dorsal surface; coxae vious in males and sometimes appears absent in females. of second legs of males with bilobate structure pos- Ventrally, carapace with small, posteriorly-projecting teroventrally, scales on lobes distally; third legs of males spines along outer margin, more numerous anterior of with paddle-like extension issuing anterodistally from respiratory areas with a few rows extending along outer the coxae, extending over bases. margin of carapace and respiratory areas to level of second legs. Respiratory areas consist of smaller, oblong Description "area" situated anterior to and abutting or nearly con- Total length (mm) 3.26-3.72 (3.53, 3.48) in males, tacting larger posterior "area"; respiratory areas distinct, 4.06-5.85 (4.92, 4.18) in females. Carapace ovoid to outlined with melanophores (Fig. 3). circular, narrower anteriorly. Carapace length (mean of Thorax dorsoventrally compressed, indistinctly seg- both sides, mm) 2.16-2.47 (2.33, 2.26) in males, 3.20- mented with two pairs of posteriorly-projecting spines 3-97 (3.57, 3-22) in females. Maximum carapace width ventrally. Spines digitate, rounded distally, the anterior (mm) 2.13-2.57 (2.41, 2.38) in males, 3.21-4.15 (3-66, pair (accessory spines) larger than posterior pair 3-30) in females. Carapace extending to middle of third (postmaxillary spines). Accessory spines situated be- legs or posterior edge of third legs/anterior margin of tween basal segments of second maxillae; postmaxillary abdomen in males. Carapace extending to middle of spines positioned farther apart than accessory spines. third legs or posterior of fourth legs/anterior of abdo- Thorax with coarse-pectinate scales scattered on ventral men in females, varies considerably in females. Females surface. Four pairs of biramous swimming legs com- -without eggs in carapace alae. Carapace in males with posed of a precoxa, coxa, basis, exopod, and endopod; inverted, dark, triangular patch of melanophores on dorsal with plumose setae on all exopods, endopods, coxae, surface of both left and right alae (much more apparent and bases. First two pairs of legs each with a recurved at lower magnifications) as well as scattered melano- flagellum on dorsal surface. Flagella laterally compressed, phores on alae and diffuse pigment scattered on cephalic bearing plumose setae. Endopods of first pair of legs region. Females with numerous scattered melanophores three-segmented, bearing three "setae" or "spines" at distal on carapace; inverted, triangular patches not as apparent end. Endopods of second pair of legs unsegmented. in females (Fig. 1). Small pores and sensilla (Fig. 2) Endopods of third and fourth pairs of swimming legs NEW SPECIES OF ARGULUS VOL. 103

FIGURES 2-3. Argulus ambystoma, new species. 2) Sensillum on dorsal surface of carapace (scale bar - 5.0 |J.m); 3) Shape of respiratory areas (right ala of female) (scale bar =1.0 mm). two-segmented. Second, third, and fourth legs of males exopods and endopods. Ventral surfaces of all leg seg- with accessory sexual structures. Coxae of male second ments, including endopod, with coarse-pectinate scales legs with a bilobate structure posteroventrally, with (Fig. 9); ventral surface of exopod with fine-pectinate scales on lobes distally (Fig. 4). Third legs each with scales (Fig. 8). Sensilla and pores scattered on ventral paddle-like extension issuing anterodistally from the surfaces of legs, pores also on dorsal surfaces of legs coxae, extending over bases (extending beyond distal (Fig. 8). margin of bases in larger males) (Fig. 4) and a hyaline, Abdomen bilobate, with pair of caudal rami near pos- bubble-like area ("socket") on posterior of bases with terior end in anal sinus. Abdomen length (mm) 1.13-1.30 opening of socket dorsally (Fig. 5). Dorsal surface of (1.21, 1.19) in males, 0.95-1.31 (1.14, 1.01) in females; basis of third leg with conical pit ("pocket") ringed with maximum width (mm) 0.94-1.06 (1.00, 0.97) in males, papillae and minute pores; posterior wall of pocket 1.07-1.40 (1.26, 1.12) in females. Anal sinus length Qun) split completely; opening of pocket leads to opening of 180-220 (202, 180) in males, 350-430 (392, 350) in fe- socket below it. Paddle-like structure of coxa extends males. Male abdomen with spine-covered crests at each over pocket. Adjacent to pocket is a small, fleshy lobe anterior corner (Fig. 10b), narrower than female abdo- tipped with papillae; lobe appears to be at base of exo- men, which is rounded at anterior corners (Fig. 10a). pod rather than on basis (Fig. 6). Bases of fourth legs Small, lateral spines along most of edge of abdomen in each with large peg, sclerotized at its base; a smaller both sexes. Each caudal ramus with five stout, naked peg (dorsal to larger peg) and small sclerotized hump "setae" (possibly not true setae) and dorsal pore just an- with minute spines ventral to larger peg (Fig. 5). Distal terior of "setae" (Figs. 11, 12). Some of the "setae" extend end of peg with notch on ventral surface and bearing beyond posterior margin of abdomen in males but not blunt "tentacles" along its rim and numerous forked in females. Paired spermathecae of female brownish, "tentacles" in concavity (Fig. 7). Precoxae + coxae of round to oval, located anteriorly on abdomen. Pair of fourth legs of males and females with posterior boot- papillae on female abdomen anteroventrally, with small shaped natatory lobes fringed with plumose setae and spines on ventral surface of papillae. Male abdomen bearing scattered, coarse-pectinate scales; distal end of with prominent black patches over testes dorsally (Fig. 1) lobes extend beyond middle of bases but not beyond and fewer, scattered melanophores ventrally over testes. distal end of bases. Dorsal surface of all legs with fine- Pores scattered on dorsal and ventral surfaces of ab- pectinate scales (Fig. 8), including dorsal surfaces of domen, but only one short, sensillum observed dorsally OHIO JOURNAL OF SCIENCE W. J. POLY 55

FIGURES 4-7. Argulus ambystoma, new species. 4) Bilobed structure (B) on posteroventral surface of coxa of second leg and paddle-like structure (P) extending from the anterodistal margin of the coxa of the third leg of male (ventral view of left, second, and third legs) (scale bar = 100.0 |Xm); 5) Socket (S) and peg (P) on third and fourth legs, respectively, of male (dorsal view) (scale bar = 50.0 (im); 6) Conical pit or "pocket" (with papillae and pores) and fleshy lobe (with papillae, indicated by arrow) located anterodorsally on basis of the third leg of male (dorsal view) (scale bar = 50.0 ]im); 7) Tip of peg on basis of fourth leg of male (dorsal view) (scale bar = 5.0 |im).

on a male's abdomen. ment fleshy, small, with few setae distally (Fig. 13). First antennae composed of four segments: first seg- Second antennae with five fleshy segments; basal seg- ment sclerotized, large, with stout posteriorly-projecting ment with posteriorly-projecting posterior spine. First posterior spine; second segment sclerotized with hump two segments rounded, bulbous; remaining three thin, anteriorly, a smaller, posteriorly-projecting medial spine cylindrical. All segments of second antennae with several and large recurved terminal spine; third segment fleshy, setae that project distally (Fig. 13). Paired postantennal cylindrical and smaller with large, stout seta distally that spines consist of reduced anterior spine and stout pos- projects ventrally and several smaller setae; fourth seg- terior spine (Fig. 13). Posterior first antennal spines and 56 NEW SPECIES OF ARGULUS VOL. 103 postantennal spines more robust in females than in First maxillae inner diameter (u.m) 290-320 (303, left: males. Reduced anterior postantennal spines vary in 300 and right: 290) (n = 15, left and right) and outer degree of development, sometimes partially hidden by diameter (|im) 430-490 (460, left: 450 and right: 450) (« = posterior first antennal spines. 15) in males, inner diameter (|lm) 500-620 (564, left: First maxillae modified into suction cups in adults. 500 and right: 500) (n = 11, left and right) and outer

FIGURES 8-10. Argulus ambystoma, new species. 8) Fine-pectinate scales on dorsal surface of exopod base; two pores are visible also (scale bar = 10.0 urn); 9) Example of a coarse-pectinate scale (on basal plate) (scale bar = 5.0 um); 10A) abdomen of female, 10B) abdomen of male, showing sexual dimorphism (dorsal view). The male abdomen has anterolateral crests with small spines, whereas the female abdomen is rounded anterolaterally and lacks the spines (scale bar 0.25 mm). OHIO JOURNAL OF SCIENCE W. J. POLY

diameter (jim) 700-850 (787, left: 700 and right: 720) (w = Table 1). Number of sclerites per support rod in males 9) in females. Number of support rods (left and right 1-5 (3-3, holotype: mean: 3.4, range: 3-4) (n = 589 rods) suction cups) in males 43-56 (49.9, left: 50 and right: 52) and in females 1-7 (4.6, allotype: mean: 4.9, range: 3-7) and in females 45-54 (49.5, left: 47 and right: 52) (see (w - 445 rods). Number of sclerites variable with position 58 NEW SPECIES OF AKGULUS VOL. 103

on suction cup, shape of sclerites variable. Usually 3 to 4 A. ambystoma from both A. maculosus and A. ameri- sclerites per rod in males, 4 to 5 sclerites per rod in fe- canus (usually 2-3 and 2 sclerites per rod in the latter males (Figs. 13, 14); lower counts, such as one, usually two species, respectively). Argulus ambystoma is similar due to missing sclerite(s), uncommon. Proximal sclerite to A. versicolor in number and shape of sclerites in usually rod- or vase-shaped, longer than other sclerites, suction cup rods, but can be distinguished readily by which may be barrel-shaped to teardrop-shaped, slightly the shape and position of the respiratory areas and pig- imbricated to offset, barely connecting with adjacent mentation of the carapace (refer to Wilson 1902, 1904; sclerites, and diminishing in size distally. Yeatman 1965 for figures of other species; except, note Second maxillae five-segmented with broad basal that at least some of the illustrations of A versicolor plate bearing three digitate spines; spines narrower, in Yeatman do not apply to that species). rounded distally. Outer two spines closer together than median spine and innermost spine; outermost spine DISCUSSION slightly shorter than remaining two spines. Basal plate Only rarely have Argulus been recorded as parasites with elevated pad bearing coarse-pectinate scales and of amphibians (for example, Goin and Ogren 1956; about three stout, simple setae that extend over base of Bower-Shore 1940; Sauer 1977; Cuvier 1798 in Wilson median posterior spine (Figs. 9, 15). Bidentate to multi- 1902; Lemos de Castro and Gomes-Correa 1985; Clark dentate spines and coarse-pectinate scales clustered in 2001). The first record of Argulus from Ambystoma distinct patches on ventral surfaces of last four seg- dumerilii was obtained when salamanders were pur- ments. Two stout, simple setae on second segment, one chased from local fishermen at Lake Patzcuaro to study on third and fourth segments posterodistally. Distal their life history (R. Brandon pers. comm.; Brandon segment with two sharp claws (usually appearing offset 1970). Salamanders harboring Argulus were captured in and not side-by-side) and blunt, elongate lobe posi- August 1968 (1 female) and December 1968 (1 female tioned above claws, short sensillum at tip of lobe. and 1 male), but the parasites were not noticed until Mouth tube short, lacking armature, not reaching tho- April and May 1969, indicating that they were too small racic accessory spines. Labium with three to four rows of to observe easily when the salamanders were initially small, embedded scales below mouth; short sensilla brought into the lab or that the species is cryptic on its scattered on remainder of labium. Pair of serrated man- host. Either may be the case, but the parasites were hidden dibles inside mouth tube. Preoral stylet present. among the gills of its host and were observed on the heads of salamanders in the evening only after the lights had Etymology been off for some time (R. Brandon pers. comm.). Also, The specific name is derived from the name of the host when the lights were turned on, the parasites quickly genus, Ambystoma (gender neuter), as a noun in apposi- moved toward the gills (R. Brandon pers. comm.). These tion to the generic name, Argulus (gender masculine). observations support the cryptic nature of this argulid on Ambystoma dumerilii and may be the reason the Remarks species was not noticed until recently. None of the speci- Argulus ambystoma is most similar to A. versicolor, mens collected by R. Brandon in 1969 could be located. A. americanus, A. maculosus, and A. diversus, but can The second record of Argulus on Ambystoma dumerilii be distinguished from them by the shape and position resulted in the collection of the material used for the of the respiratory areas and the secondary sexual modi- description; R. Brandon received some A. dumerilii from fications on the legs of males. The paddle-like structure the Cincinnati Zoo and collected the Argulus from these on the third leg of male Argulus ambystoma resembles . An additional observation of Argulus around the those of males of A. americanus and A. diversus. The gills and on the head of wild caught A. dumerilii was number of suction cup rod sclerites also distinguishes supplied by S. Randal Voss (pers. comm.) in 1996.

TABLE 1

First maxillae support rod counts for male and female Argulus ambystoma, new species (holotype and allotype counts in bold).

Male (n = 7) Female (n = 6)

Left Suction Cup 49 47 46 48 50 53 53 53 47 45 49 52 46

Right Suction Cup 48 51 43 50 52 52 56 48 52 -a 49 50 54

Mean = 49.9; Range = 43-56 Mean = 49.5; Range = 45-54

Mean = 49.7; Range = 43-56 (male and female)

"Structure missing. OHIO JOURNAL OF SCIENCE W. J. POLY 59

The number of suction cup rods is a useful taxo- because one seta is often hidden by the other two setae. nomic character that rarely has been included in de- Argulus ambystoma, A. americanus, A. maculosus, A. scriptions or redescriptions. A neglected aspect of versicolor, A. diversus, A. stizostethii, A. flavescens, A. suction cup sclerites has been variability within a species foliaceus, A. meehani Cressey, and A. chesapeakensis and on a single suction cup. Rizvi (1970) examined Cressey all have five "setae" on the caudal rami (this suction cup rods of Argulus foliaceus (Linne), and rod study; Shimura and Asai 1984; Yeatman 1965; Kellicott number was 40 to 51 (10 specimens). Argulus japoni- 1880; Suarez-Morales and others 1998; Leydig 1889; cus suction cup rod numbers varied from 38-50 with Cressey 1971), whereas A. ellipticaudatus Wang and A. a mean of 44.6 (12 specimens), and asymmetry (left/right) melanostictus apparently have only four (Wang I960; was observed in all 12 individuals (Hsiao 1950). Pilgrim Benz and others 1995), A. foliaceus (3rd stage) has four (1967) examined A, japonicus from New Zealand and (Rushton-Mellor and Boxshall 1994), and A. japonicus st reported a range of 44-50 suction cup rods with a mean (1 stage) has three (Tokioka 1936). of 45.9 (4 specimens, 7 suction cups). Suction cup rods The pocket on the third leg of male A. ambystoma of 15 Argulus matuii Sikama (29 suction cups) ranged is similar to the pocket of male A. appendiculosus from 75-83 in males and 81-95 in females with sym- Wilson shown by Sutherland and Wittrock (1986). Male metry occurring in only three individuals (Sikama 1938). A. versicolor'And A. americanus also have pockets similar Argulus ambystoma suction cup rods ranged from 43 to those of the above-mentioned species (pers. observ.). to 56 with a mean of 49.7 (13 specimens, 25 suction Sensilla of Argulus rarely have been mentioned or cups), and only one individual was not asymmetrical figured in the literature. Leydig (1889) and Debaisieux (Table 1). Variation in both sclerite number and shape (1953) illustrated the type of sensillum shown herein has been noted by a few authors, depending on the (Fig. 2) as well as other sensilla found on A. foliaceus. position of rods in a suction cup (Fryer 1959; Rushton- Madsen (1964:22-23) mentioned the sensilla ("hairs") Mellor and Boxshall 1994; Avenant-Oldewage and found on the rim of the carapace, anterior rim of head, Oldewage 1995). Variation in sclerite number and shape and dorsal surface of carapace and stated that the also occurs in A. ambystoma and other North American sensilla were probably rheotactical. Smaller sensilla are Argulus spp. (pers. observ.). In A. ambystoma higher abundant in the interspaces between the larger sensilla numbers of sclerites per rod usually occur in the antero- and have been figured most often on nauplius stages lateral (outer) section of a suction cup rim, and the (Tokioka 1936; Rushton-Mellor and Boxshall 1994). sclerites tend to be more bulbous or round antero- Linnenbach and Hausmann (1983) included a photo- laterally, whereas sclerite numbers are lower posteriorly micrograph of sensilla on Argulus sp., and sensilla and and on the inner margin, and sclerites tend to be more various scale types of A. foliaceus were shown in SEM rod-like and slender; Fryer (1959) found the same trend micrographs by Lange and Sundermann (1990). Suther- land and Wittrock (1986:410) reported only sensory pits in Argulus ambloplites Wilson. One female A. amby- on the dorsal surface of the carapace of A. appen- stoma had one long, unsegmented rod and a two-seg- diculosus, hence there may be some taxonomic value mented rod, both spanning the entire width of the in the distribution or presence/absence of sensilla as suction cup rim. Intraspecific variation in the length of shown among the Copepoda (for example, Fleminger the carapace alae also can be found among Argulus 1973). No pores or sensilla were observed on the dorsal spp. (Meehean 1940; Fryer 1982), and A. ambystoma surface of the thorax of A. ambystoma in this study. exhibited such variation. Number of setae (or "spines") on the endopod of the The dorsal pores on the caudal rami have not been first leg and number of "setae" on the caudal rami may reported previously in the genus Argulus nor in any be useful characters for or systematics. Sikama other member of the Branchiura, but similar pores on the ventral surface of the caudal rami of Unicolax col- (1938) indicated that three "spines" were present at the lateralis Cressey and Boyle Cressey (Copepoda: Bom- tip of the endopod in Argulus matuii (see his Fig. 11). olochidae) were shown by Cressey and Boyle Cressey Rushton-Mellor and Boxshall (1994) indicated that two (1980, their Fig. 113e); however, they did not mention setae were present on the last segment of the endopod the pore specifically. The function of these pores is un- of the first leg of Argulus foliaceus in the first develop- known. A small cyst measuring 160 Lim was extracted mental stage, whereas three setae were present in all from the left ala of the carapace of the holotype, but its later stages. Argulus japonicus has three setae at the tip identity has not been determined; another cyst may of the endopod of the first leg, and three setae are exist in the coxa of the right, second leg of a female present in the first naupliar stage according to Tokioka individual, but no attempt was made to extract or posi- (1936). Argulus stizostethii Kellicott, A. flavescens, A. tively identify the object. Lesions characteristic of crus- rhipidiophorus Monod, and A. major Wang also have tacean shell disease were present on two males and the three setae (Kellicott 1880; Suarez-Morales and others three females. 1998; Monod 1931; Wang I960); Argulus ambystoma possesses the three setae as well, at least in adults. Benz ACKNOWLEDGMENTS. Joseph A. Beatty (Southern Illinois University [SIU]) and others (1995) stated that two spines were present provided specimens of Argulus ambystoma for the description, and at the tip of the endopod of A. melanostictus, as did Ronald A. Brandon (SIU) and S. Randal Voss (University of California, Dana and Herrick (1837) for A. catostomi Dana and Davis) supplied observations of Argulus on Ambystoma dumerilii. Randall Tindall, Steven Schmitt, John J. Bozzola, and Dee Gates (SIU, Herrick, but there are three setae in these species as Integrated Microscopy and Graphics Expertise [I.M.A.G.E.]) contributed well (pers. observ.). Three setae are difficult to observe their knowledge of specimen preparation and SEM techniques and 60 NEW SPECIES OF ARGULUS VOL. 103

helped on many occasions with specimen preparation, examination, freshwater fishes: a handbook and key. Freshwater Biol Assn and photoaraphy, and Cheryl Broadie and Steve Mueller (SIU, Scientific Publ No. 46, Cumbria, UK. 87 p. I.M.A.G.E.) prepared the figures. Ardis Johnston (Museum of Com- Fucugauchi Suarez del Real MG, Garcia Magana L, Brito Arjona B parative Zoology) and Raymond B. Manning (deceased), Paula del R. 1988. Analisis previo de la parasitofauna de peces de la Rothman, Janice Walker, Karen Reed, and Chad Walter (National Laguna del Rosario, Huimanguillo, Tabasco. Divulgacion Cientifica Museum of Natural History) kindly loaned specimens for comparative (Diciembre 1988):319-35. studies and cataloged the A. ambystoma specimens. The science Goin CJ, Ogren LH. 1956. Parasitic copepods (Argulidae) on amphib- librarians at Morris Library (SIU) were instrumental in obtaining much ians. J Parasitology 42:172. of the relevant literature. 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